The millipede family Paradoxosomatidae in Paraguay, with descriptions of five new species (Diplopoda, Polydesmida)

Revue suisse de Zoologie 112 (4): 807-830; decembre 2005

The millipede family Paradoxosomatidae in Paraguay, with
descriptions of five new species (Diplopoda, Polydesmida)

Sergei I. GOLOVATCH

Institute for Problems of Ecology and Evolution, Russian Academy of Sciences,
Leninsky pr. 33, Moscow 119071 (V-71), Russia. E-mail: sgol@orc.ru

The millipede family Paradoxosomatidae in Paraguay, with descrip-
tions of five new species (Diplopoda, Polydesmida). - A review is pro-
vided of the millipede family Paradoxosomatidae in Paraguay, with keys
compiled to 14 genera and 63 species occurring in this country and/or
adjacent areas. Five species are described as new: Catharosoma bilineatum
sp. n., C. mahnerti sp. n., Broelemannopus minutus sp. n., Mestosoma sim-
plex sp. n. and M. crassipes sp. n.

Keywords: Diplopoda - Polydesmida - Paradoxosomatidae - taxonomy -
Paraguay.

INTRODUCTION

The family Paradoxosomatidae is among the largest among the Diplopoda, with
nearly 200 genera currently accepted as valid. The American fauna is highly peculiar
due to the absence of indigenous paradoxosomatids north of Costa Rica. In South
America two major centres of diversification are distinguishable, one in the area of
southern Brazil, Paraguay and northern Argentina, the other in Peru, northern Bolivia
and possibly Ecuador. The vast regions of northern Brazil, Guiana, Venezuela and
Colombia appear to harbour very few species (Jeekel, 1968, 2002).

The Neotropical fauna itself is strongly dominated by relatively few (about a
dozen) genera of the endemic tribe Catharosomatini. Of these genera, Mestosoma
Silvestri, 1897 is certainly the largest and particularly widespread, with about 75
species occurring throughout South America and reaching both Costa Rica and the
island of Dominica (introduced?) in the north. Catharosoma Silvestri, 1913 is the
second largest genus, its 13 species being known from Brazil, Paraguay and Argentina
(Hoffman, 1980, 1999). The remaining genera are mono- to oligotypic.

The present paper provides a review of, and keys to, the bulk of the paradoxo-
somatids occurring in the southern Neotropical diversification centre, based both on all
available literature sources and on the important collection of Paradoxosomatidae from
Paraguay housed in the Museum d’histoire naturelle, Geneva (MHNG). Five new
species from three genera of Catharosomatini have been revealed there, all described
below and thus considerably enriching our knowledge of the fauna of Paraguay. A few
paratypes from the material have been retained for the collection of the Zoological
Museum, State University of Moscow (ZMUM), Russia.

Manuscript accepted 07.02.2005

808

S. I. GOLOVATCH

FAUNISTIC REVIEW

Since detailed reviews of the history of research on Neotropical Paradoxo-
somatidae are available and still fully relevant (Jeekel, 1963, 1968), only a short
account appears necessary here concerning the species list of the particular region
involved. Virtually all later contributions (e.g. Hoffman, 1977, 1999; Golovatch, 1992;
Jeekel, 2002; Golovatch et al ., 2003) are irrelevant in the present context, as they only
or chiefly treat more northern faunas.

The following Paradoxosomatidae from Paraguay and/or the adjacent parts of
Bolivia, northern Argentina and southern Brazil have hitherto been recognized as valid:

Broelemannopus Verhoeff, 1938

Broelemannopus escaramucensis (Schubart, 1944) - Brazil (Sao Paulo)
(Schubart, 1944, 1952)

B. glabratus (Schubart, 1945) - Brazil (Rio de Janeiro) (Schubart, 1945b)

B. ibitiensis (Schubart, 1945) - Brazil (Sao Paulo) (Schubart, 1945a)

B. pirassunungensis (Schubart, 1944) - Brazil (Sao Paulo) (Schubart, 1944,
1945a, 1952)

Catharosoma Silvestri, 1897

Catharosoma curitibense Schubart, 1953 - Brazil (Parana) (Schubart, 1953)

C. digitale Schubart, 1953 - Brazil (Parana) (Schubart, 1953)

C. hoffinani Kraus, 1956 - Paraguay (Kraus, 1956)

C. intermedium (Carl, 1902) - Brazil (Rio Grande do Sul) (Carl, 1902; Attems,
1914, 1937)

C. mesorphinum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914,
1937; Schubart, 1953)

C. mesoxanthum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914,
1937; Schubart, 1953)

C. mixtum Kraus, 1956 - Brazil (Santa Catarina) (Kraus, 1956)

C. mynnekurum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914,
1937; Schubart, 1953)

C.palmatum Schubart, 1953 - Brazil (Parana) (Schubart, 1953)

C.palustre Schubart, 1943 - Brazil (Mato Grosso) (Schubart, 1943)

C. paragnayense (Silvestri, 1895) - Paraguay, Argentina and Brazil (Santa
Catarina and Mato Grosso) (Silvestri, 1895b, 1902; Attems, 1898, 1914, 1937;
Schubart, 1953; Kraus, 1956; Jeekel, 1965) («South America», according to Mauries,
1998)

C. peraccae Silvestri, 1902 - Paraguay (Silvestri, 1902; Attems, 1914, 1937)

C. taeniatum (Brolemann, 1929) - Brazil (Santa Catarina) (Brolemann, 1929;
Attems, 1937; Schubart, 1953)

Chondromorpha Silvestri, 1897

Chondromorpha xanthotricha (Attems, 1898) - pantropical, introduced (e.g.
Jeekel, 1963)

PARADOXOSOMATIDAE IN PARAGUAY

809

Gonodrepanoides Schubart, 1945

Gonodrepanoides travassosi Schubart, 1945 - Brazil (Rio de Janeiro)
(Schubart, 1945b)

Gonodrepanum Attems, 1914

Gonodrepanum drepanephoron (Attems, 1898) - Brazil (Sao Paulo) and
Argentina (Attems, 1898, 1901, 1914, 1937; Brolemann, 1902b; Mauries, 1998)
G.falciferum Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b)

G . flavolineatum Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b)
G.furcatum Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b)

G. grajahuense Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b)

G. levisetum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914,
1937; Schubart, 1953; Mauries, 1998)

G. levisetum var. coniferum (Attems, 1898) - Brazil (Santa Catarina) (Attems,
1898, 1914, 1937; Schubart, 1953; Mauries, 1998)

G. torresae Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b)

Habrodesmoides Attems, 1943

Hcibrodesmoides costcilimcii (Schubart, 1945) - Brazil (Rio de Janeiro)
(Schubart, 1945b)

H. perturbans Attems, 1943 - Brazil (Rio de Janeiro) (Attems, 1943)
Mestosoma Silvestri, 1897

Mestosoma alticola (Attems, 1931) - Bolivia (Cochabamba) and Peru (Attems,
1931, 1937; Kraus, 1956)

M. bcilzcinii (Silvestri, 1895) - Bolivia (Yungas) (Silvestri, 1895a; Attems, 1914,

1937)

M. bicolor Silvestri, 1898 - Paraguay and Brazil (Mato Grosso) (Silvestri, 1898,
1902; Attems, 1899, 1914, 1937; Jeekel, 1965)

M. boliviae (Chamberlin, 1957) - Bolivia (near Potosi) (Chamberlin, 1957)

M. borellii (Silvestri, 1895) - Argentina (Tucuman) and Paraguay (Silvestri,
1895b; Attems, 1914, 1937; Mauries, 1998)

M. camerani (Silvestri, 1895) - Argentina (Chaco) (Silvestri, 1895b; Attems,
1914, 1937; Jeekel, 1965; Mauries, 1998)

M. carioca (Schubart, 1945) - Brazil (Rio de Janeiro) (Schubart, 1945b)

M. dereliction (Silvestri, 1895) - Bolivia (Misiones Mosetenes) (Silvestri,
1895a; Attems, 1914, 1937)

M. dijferens Kraus, 1956 - Bolivia (Cochabamba) (Kraus, 1956)

M.femorale (Schubart, 1943) - Brazil (Mato Grosso) (Schubart, 1943)

M. kalliston (Attems, 1898) - Brazil (Rio Grande do Sul) (Attems, 1898, 1914,

1937)

M. luctuosum Silvestri, 1897 - Bolivia (Caiza) (Silvestri, 1897; Attems, 1914,

1937)

810

S. I. GOLOVATCH

M. lugubre Silvestri, 1897 - Argentina (Buenos Aires) and Uruguay (Silvestri,
1897, 1902; Attems, 1914, 1937; Mauries, 1998)

M. montanum (Silvestri, 1895) - Bolivia (Yungas) (Silvestri, 1895a; Attems,
1914, 1937)

M.perfidum (Schubart, 1943) - Brazil (Sao Paulo) (Schubart, 1943)

M. pseudomorphum (Silvestri, 1895) - Paraguay (Silvestri, 1895b; Carl, 1902;
Attems, 1914, 1937; Jeekel, 1965)

M . pulvillatum (Attems, 1898) - Paraguay (Attems, 1898, 1901, 1914, 1937)
M. salvadorii (Silvestri, 1895) - Argentina (Salta), Bolivia (Chaco) and
Paraguay (Silvestri, 1895b, 1902; Attems, 1914, 1937; Hoffman, 1977; also Costa
Rica, according to Mauries, 1998)

M. schindleri Kraus, 1956 - Bolivia (Silhuencas) (Kraus, 1956)

M. tricuspis (Verhoeff, 1938) - Paraguay (Verhoeff, 1938)

M. truncation (Schubart, 1943) - Brazil (Mato Grosso) (Schubart, 1943)

M. vittatum (Attems, 1898) - Paraguay (Attems, 1898, 1914, 1937)

Mogyella Schubart, 1944

Mogyella nana Schubart, 1944 - Brazil (Sao Paulo) (Schubart, 1944, 1952)
Mogyosoma Schubart, 1944

Mogyosoma hamatum Schubart, 1944 - Brazil (Sao Paulo) (Schubart, 1944)
Ologonosoma Silvestri, 1898

Ologonosoma iguassuense (Schubart, 1953) - Brazil (Parana) (Schubart, 1953;
Mauries, 1998)

O. sanctum (Silvestri, 1895) - Paraguay (Silvestri, 1895a; Attems, 1914, 1937;
Mauries, 1998)

Orthomorpha Bollman, 1893

Orthomorpha coarctata (De Saussure, 1860) - pantropical, introduced (e.g.
Jeekel, 1963)

Oxidus Cook, 1911

Oxidus gracilis (C. L. Koch, 1847) - subcosmopolitan, introduced (e.g. Jeekel,

1963)

Promestosoma Silvestri, 1898

Promestosoma boggianii Silvestri, 1898 - Paraguay (Silvestri, 1898; Attems,
1899, 1914, 1937; Jeekel, 1965; Mauries, 1998) and Brazil (Mato Grosso and Mato
Grosso do Sul) (original data)

Pseudogonodrepanum Schubart, 1945

Pseudogonodrepanum scitum Schubart, 1945 - Brazil (Rio de Janeiro)
(Schubart, 1945b)

PARADOXOSOMATIDAE IN PARAGUAY

811

A few more species, i.e. Mestosoma laetum Silvestri, 1897 (Bolivia, San
Francisco - Silvestri, 1897), «Strongylosoma» nitidum Brolemann, 1902 (Brazil,
Cubatao, Sao Paulo - Brolemann, 1902a), «Strongylosoma» pustulatum Brolemann,
1902 (Brazil, Sao Paulo - Brolemann, 1902b) and Catharosoma bromelicola Schubart,
1945 (Brazil, Rio de Janeiro - Schubart, 1945b), the male characters of which are not
or are insufficiently known, still remain incertae sedis\ none can be included in a key
(Jeekel, 1963).

The same concerns lulidesmus Silvestri, 1895, with its type-species I. typicus
Silvestri, 1895 known only from a female holotype from Bolivia (Yungas) (Silvestri,
1895a). Jeekel (1963) correctly excluded lulidesmus from Paradoxosomatidae on the
basis of a later redescription of I. typicus from Chilean material provided by Silvestri
(1905). But Hoffman (1980), rightly doubting the conspecificity of the Bolivian and
Chilean samples, believes that the holotype of I. typicus represents a Mestosoma. If so,
then Mestosoma becomes a junior synonym of lulidesmus. Only topotypic material in
connection with revisionary work can shed additional light on the identity of all of the
enigmatic taxa mentioned above.

DESCRIPTIONS OF NEW SPECIES

Catharosoma bilineatum sp. n. Figs 1-9

Material: Holotype 8 (MHNG), Paraguay, Alto Parana Prov., Forestry Centre
(C.F.A.P.), Puerto Presidente Stroessner, plot 7, 18.03.1983, leg. P. Berner & C. Dlouhy. -
Paratypes: 1 9 (MHNG), same locality, together with holotype; 1 <3,3 9 9 (MHNG), 1 d, 1 9
(ZMUM), same locality, around Pindo trunk, plot 3, 24.03.1983, leg. C. Dlouhy; 1 9 (MHNG),
same locality, soil sample, plot 4 (Monte natural), Winkler extraction, 10.03.1983, leg. C.
Dlouhy; 1 9 (MHNG), same plot 4 (Monte natural), 10.03.1983, leg. P. Berner & C. Dlouhy; 1
9 (MHNG), same locality, soil sample, plot 2 (Monte natural), 29.02.1983, leg. C. Dlouhy; 1 8
(MHNG), same locality, soil sample, plot 8 (Monte natural), 18.03.1983, leg. P. Berner & C.
Dlouhy.

Name: To emphasize the presence of two dark paramedian stripes divided by a similar-
ly wide but light axial stripe.

Diagnosis : Differs from congeners by the characteristic coloration, the strong-
ly reduced paraterga visible only on segments 2-4, the presence of relatively short an-
tennae, the deeply divided sternal lobe present between coxae 6 of the 8 , the subunci-
form tip of the hypoproct, the bifid tip of the epiproct, coupled with certain details of
solenophore structure.

Description : Length 21-27 mm (<?, 9), width of midbody metazona 1 .9-2.1
(<3) or 2. 1-2. 8 mm (9); 9 usually larger and somewhat broader than 8 . Holotype ca
23 mm long and 2.0 mm wide. Coloration in alcohol pale yellowish brown to brown,
with a characteristic pair of brown to dark brown paramedian stripes divided by an
equally wide axial stripe of background coloration; legs and sterna yellowish gray,
slightly paler than background coloration; tip of antennae pallid, penultimate anten-
nomere dark brown; posterior half of body always somewhat paler than anterior one.

Body subcylindrical, not moniliform. Postcollar constriction faint, width of
head = 5-16 > collum = 4 > 2 = 3; on segments 17-20 trunk gradually and gently ta-
pering toward telson both in width and in height. Antennae relatively short, slightly
clavate, in situ reaching beyond segment 2 dorsally (Fig. 1), a little shorter in 9.

812

S. I. GOLOVATCH

Catharosoma bilineatum sp. n., 6 holotype: 1) anterior body portion, lateral view; 2) telson, lat-
eral view; 3) sternal structures between coxae 4-7 (coxae 4 on top); 4) sternal structures between
midbody coxae; 5) leg 12; 6-9) left gonopod, medial, ventral, lateral and ventromedial views, re-
spectively. Scale bars 1.0 (1-5) and 0.25 mm (6-9).

Paraterga strongly reduced, only on segment 2 fully developed as low keels with a
caudal tooth, discernible on segments 3 and 4 as arcuated sulci (Fig. 1), on following
segments totally missing. Tegument smooth, shining, only rear parts of metazona often
faintly rugulose. Limbus thin, caudal margin entire. Metaterga fully devoid of a trans-
verse sulcus; setae medium-sized, often abraded, arranged 2+2 in a transverse line
behind a shallow stricture between pro- and metazona. Ozopores lateral, opening level
to metatergal surface ca 1/3 metazonital length away from caudal edge. Pleurostemal
carinae shaped like narrow keels with a caudal spinule (Fig. 1) traceable until segment
16(2) or 17 (d), on following segments wanting. Epiproct (Fig. 2) rather long, coni-
form, bifid, in 3 a little longer than in 2 . Hypoproct (Fig. 2) subtriangular, tip pointed,
unciform, directed ventrad; 1+1 paramedian setae at caudal edge considerably
separated from each other. Paraterga, pleurostemal carinae and hypoproct uncus in <5
a little more strongly developed than in 2 .

Sterna densely setose. Sternum between coxae 3 of 6 with a paramedian, nearly
contiguous pair of bunches of setae. Sternum between coxae 4 of 6 with a prominent,
linguiform, laterally setose outgrowth directed an tero ventrad (Fig. 3); a similar but

PARADOXOSOMATIDAE IN PARAGUAY

813

much lower outgrowth between coxae 5 of S (Fig. 3); a more or less prominent and
deeply divided outgrowth between coxae 6 of S poorly delimited caudally against a
similar but less prominent bulge with peculiar paramedian bunches of setae between
coxae 7 of S (Fig. 3). Postgonopodial sterna with shorter (between anterior coxae) or
longer (between posterior coxae), coniform, paramedian spines developed a little bet-
ter in S (Fig. 4) than in 2 .

Legs without tarsal brushes but densely setose ventrally; setation gradually
thinning out toward telson; tibiae swollen ventrally and supporting characteristic
brushes (Fig. 5) between leg-pair 9 until two last pairs. Legs in 8 a little longer than
in 2 , as usual becoming a little longer and slenderer toward telson. Each coxa 2 of 6
with a prominent, somewhat sinuate, disto ventral spine carrying a gonopore at base.

Gonopods (Figs 6-9) highly complex. Coxite elongate, subcylindrical, setose
disto ventrally; cannula normal. Telopodite strongly unciform, with a hypertrophied, as
usual densely setose prefemoral part somewhat longer than femorite. A few undu-
lations on ventrocaudal face and a few longitudinal ridges anterodorsally at base of
solenophore. Solenophore coiled, ventrally with both a slightly folded base of lamina
medialis (m) and a subtriangular parabasal lobe (p) similarly well-developed; p
followed by a spine (k) supporting the tip of a flagelliform solenomere at base of a
hyaline lobule (h), this lobule supporting the penultimate 1/4 extent of both lamina
lateralis and lamina medialis; distalmost 1/4 extent of both laminae slender, unciform,
free of support.

Remarks: Judging from the colour pattern and gonopod structure, this new
species seems to be especially close to C. mesoxanthum , but both differ in size, in
structure of the process between coxae 6 of the <?, and in direction of the apical
papillae on the epiproct (see also key below).

Catharosoma mahnerti sp. n. Figs 10-17

Material: Holotype S (MHNG), Paraguay, Prov. Concepcion, Estancia Laguna Negra,
15 km E of Paso Barreto, dead wood, 13.10.1985, leg. Expedition zoologique du Museum de
Geneve. - Paratypes: 1 6 , 2 2 2 (MHNG), 1 <3, 1 2 (ZMUM), same locality, together with
holotype; 1 S (MHNG), Prov. Concepcion, near mouth of Ao River, Trementina (Estancia
Laguna Negra), forest, 13.10.1985, leg. Expedition zoologique du Museum de Geneve.

Name: Honours Dr Volker Mahnert, who entrusted me this valuable material for study.

Diagnosis: Differs from congeners by the submoniliform body, the longer
antennae, the absence of a distinct colour pattern and of an unciform hypoproct, the
presence of two paramedian pilose ridges between coxae 3 of the 6 , and in certain
details of gonopod structure.

Description: Length 20-23 mm (6\ 2), width of midbody metazona 1.7 (c?) to
2.3-2.5 mm (2); 2 usually larger and somewhat broader than 6 . Holotype ca 20 mm
long and 1 .7 mm wide. Coloration in alcohol uniformly light brown to red-brown; only
penultimate antennomere rather dark brown, tip of antenna pallid; legs paler, yellow-
brown to brown.

Body subcylindrical, submoniliform. Postcollar constriction faint, width of
head = 5-16 > collum = 4 > 2 = 3; on segments 17-20 trunk gradually and gently
tapering toward telson both in width and in height. Antennae medium-sized, slender, in
situ reaching beyond segment 3 dorsally (Fig. 10), a little shorter in 2. Paraterga

814

S. I. GOLOVATCH

Figs 10-17

Catharosoma mahnerti sp. n., 6 paratype: 10) anterior body portion, lateral view; 11) metater-
gum 10, dorsal view; 12) telson, lateral view; 13) hypoproct, ventral view; 14) sternal structures
between coxae 2-7 (legs 2 at left); 15) sternal structures between midbody coxae; 16) leg 15; 17)
right gonopod, medial view. Scale bars 1.0 (10-16) and 0.2 mm (17).

strongly reduced, only on segment 2 fully developed as low keels devoid of a caudal
tooth, discernible on segments 3 and 4 as arcuated lines, on segment 5 as modest
swellings (Fig. 10), on following segments totally missing. Tegument smooth, shining,
only metazona at places faintly rugulose. Limbus thin, caudal margin entire. Metaterga
fully devoid of a transverse sulcus; setae short, often abraded, arranged 3+3 on seg-
ments 2 and 3, on following segmens 2+2 in a transverse line behind a rather deep
stricture between pro- and metazona (Fig. 11). Ozopores lateral, opening level to
metatergal surface ca 1/3 metazonital length away from caudal edge (Fig. 10).
Pleurostemal carinae like narrow keels with a caudal spinule (Fig. 10) traceable until
segment 16 (9) or 17-18 (cJ), onward wanting. Epiproct (Fig. 12) rather long, coni-
form, faintly emarginate at tip in dorsal view, in 6 a little longer than in 9 . Hypoproct

PARADOXOSOMATIDAE IN PARAGUAY

815

(Figs 12, 13) subtriangular, tip narrowly rounded, nearly pointed, straight; 1+1 para-
median setae at caudal edge poorly separated. Paraterga, pleurostemal carinae and
hypoproct a little more strongly developed in S than in 2 .

Sternal structure (Figs 14, 15) much like in C. bilineatum sp. n., but with two
paramedian pilose ridges between coxae 3 of 8 and paramedian bunches of setae on
poorly developed knobs between coxae 6 of 8 .

Legs without tarsal brushes but densely setose ventrally; setation gradually
thinning out toward telson; tibiae swollen ventrally and supporting characteristic
brushes (Fig. 16) between leg-pair 7 and two last pairs. Legs in 8 a little longer than
in 2 , as usual becoming a little longer and slenderer toward telson. Each coxa 2 of <5
with a strong, distoventral, sigmoid, apically pointed process carrying a gonopore at
base (Fig. 14).

Gonopods (Fig. 17) highly complex, much like in C. bilineatum n. sp., but cox-
ite more elongate, telopodite circular and elongate, while solenophore ventrally with a
much larger, hyaline lobe/base of lamina medialis (m), an apically strongly unciform
parabasal lobe (p) followed by a spiniform process supporting a flagelliform
solenomere at base of a hyaline lobule (h), this lobule supporting the penultimate 1/4
extent of both lamina lateralis and lamina medialis; distalmost 1/4 extent of both
laminae likewise slender, unciform, free of support.

Remarks: Judging from the particularly elongate gonopod telopodite, which
includes the femorite, this new species seems to be especially close to C. curitihense
and C. intermedium , but the circular telopodite and the shape of its individual elements
in C. mahnerti sp. n. are quite characteristic.

At first Jeekel (1963) merged Catharosoma and Mestosoma together, but later
(1968) he separated them on the account of the apparently more apomorphic character
states observed in Mestosoma species. This viewpoint has also been shared by
Hoffman (1980).

Broelemannopus minutus sp. n. Figs 18-25

Material: Holotype 8 (MHNG), Paraguay, Alto Parana Prov., Forestry School, 12 km S
of Puerto Presidente Stroessner, soil sample 1 , 04.1983, leg. C. Dlouhy. - Paratypes: 20 8 8 ,22
2 2,2 juv. (MHNG), 2 8 8, 2 2 2 (ZMUM), same locality, together with holotype; 1 8 , 1 2
(MHNG), same locality, soil sample 8, 04.1984; 2 8 8 , 1 2 (MHNG), same locality, soil sample
24, 06.1984; 3 8 8 (MHNG), same locality, soil sample 10,05.1984; 1 2 , 3 juv. (MHNG), same
locality, soil sample 22, 25.06.1984; 1 8, 1 2 (MHNG), same locality, soil sample 25,
25.06.1984; 1 8 (MHNG), same locality, soil sample 7,04.1984; 1 8, 3 juv. (MHNG), same lo-
cality, soil sample 23, 25.06.1984; 2 8 8 (MHNG), same locality, soil sample 3, 21.05.1984; 1
8,1 2,1 juv. (MHNG), same locality, soil sample 13, 04.1984; 1 <3,1 2,1 juv. (ZMUM), same
locality, soil sample 30, 04.1984; 14 88, 17 2 2 (MHNG), same locality, soil sample 12,
04.06.1984, all leg. C. Dlouhy.

Name: To emphasize the small body size.

Diagnosis: Differs from congeners by the particularly small size, the especially
strongly reduced paraterga visible only on segment 2, the lack of transverse metatergal
sulci, the absence of a distinct colour pattern and of an unciform hypoproct, and in
certain details of solenophore structure.

Description: Length 7-9 mm (<?, 2), width of midbody metazona 0.6-0 .7 (<3)
to 0.8-0 .9 mm ( 2 ); 2 usually larger and somewhat broader than 8 . Holotype ca 8 mm

816

S. I. GOLOVATCH

Figs 18-25

Broelemannopus minutus sp. n., 6 paratype: 18) anterior body portion, lateral view; 19) caudal
body portion, lateral view; 20) sternal structures between coxae 3-7 (coxae 3 on top); 21) ster-
nal structures between midbody coxae; 22) leg 10; 23-25) right gonopod, medial, lateral and an-
terodorsal views, respectively. Scale bars 0.5 (18-19), 0.25 (20-22) and 0.1 mm (23-25).

long and 0.7 mm wide. Coloration in alcohol uniformly brown to red-brown, only
juveniles, antennae and legs paler, yellow-brown to brown; tip of antenna pallid.

Body subcylindrical, not moniliform. Postcollar constriction faint, width of
head > collum > 2 > 3 < 4 < 5-16; on segments 17-20 trunk gradually and gently
tapering toward telson both in width and in height. Antennae medium-sized, slender, in
situ reaching the end of segment 2 dorsally (Fig. 18), a little shorter in 2 . Paraterga
strongly reduced, only present as low lines devoid of a caudal tooth on segment 2
(Fig. 18), on following segments totally missing. Tegument smooth, shining. Limbus
thin, caudal margin entire. Metaterga fully devoid of a transverse sulcus; setae
relatively long, sometimes abraded, arranged 2+2 on segment 2, on following seg-
ments 1+1 in a transverse line behind a shallow stricture between pro- and metazona
(Fig. 18). Ozopores lateral, opening level to metatergal surface ca 1/3 metazonital
length away from caudal edge. Pleurostemal carinae like narrow arcuated lines (Fig.
18) traceable until segment 16(2) or 17 (cJ), on following segments wanting. Epiproct
(Fig. 19) rather long, coniform, in £ a little longer than in 2 , tip with a stong claw-
shaped uncus directed dorsocaudally. Hypoproct (Fig. 19) subtri angular, tip narrowly
rounded, nearly pointed, straight; 1+1 very long paramedian setae at caudal edge
poorly separated from each other. Paraterga. pleurostemal carinae, epi- and hypoproct
in 6 a little more strongly developed than in 2 .

PARADOXOSOMATIDAE IN PARAGUAY

817

Sterna mostly sparsely setose. Sternum between coxae 3 of 6 furnished with
several transverse rows of relatively small, subcontiguous, scale-like structures on a
boss (Fig. 20). Sterna between coxae 4 and 5 of 6 with a very prominent, linguiform,
laterally setose outgrowth directed anteroventrad (Fig. 20); a paramadian pair of
bunches of setae between coxae 5 and 7 of c? (Fig. 20); a central, compact and thicker
bunch of setae on a swelling between coxae 6 of 8 (Fig. 20). Postgonopodial sterna
without modifications (Fig. 21).

Legs without tarsal brushes but densely setose ventrally, without modifications
(Fig. 22); setation gradually thinning out toward telson; legs in 6 a little longer than
in 9 , as usual becoming a little longer and slenderer toward telson. Each coxa 2 of 6
nearly unmodified, with a small distoventral cone surmounted by a gonopore.

Gonopods (Figs 23-25) not particularly complex. Solenophore ventrally with a
distinct but not hypertrophied, rounded, hyaline lobe/base of lamina medialis (m);
another simple, parabasal lobe (p) supporting the tip of a flagelliform solenomere at
base of a smaller hyaline lobule (h), the latter supporting the subterminal part of both
lamina lateralis and lamina medialis; distalmost extent of both laminae very short and
rather broad, subunciform and subtruncate.

Remarks: Judging from the particularly small size, the similarly uncigerous
epiproct, the paraterga present on segment 2 only, as well as from several other traits,
including those of the gonopod, the new species seems to be especially close to
B. ibitiensis , from Monte Alegre, Sao Paulo State, Brazil (Schubart, 1945a). The
distinctions of B. minutus lie in the total absence of sternal cones on postgonopodial
segments of the 6 , as well as the presence of a smaller and differently armed lobe
between coxae 3 of the 6 , and of a smaller lobe p on the solenophore (see also key
below). B. ibitiensis has only tentatively been placed in Broelemannopus (see Jeekel,
1963) but, since this species shows especially close affinities with B . minutus sp. n.,
there can no longer be any doubt that both are congeners best to be assigned to
Broelemannopus .

In general the genus Broelemannopus is so poorly distinguished from
Mestosoma that at first Jeekel (1963) merged these two genera together and treated the
former as only a species group of the latter. However, later (1968) he separated them
on the basis of certain apomorphies observed in both (see key below), a view which is
also shared by Hoffman (1980).

Mestosoma simplex sp. n. Figs 26-32

Material: Holotype 6 (MHNG), Paraguay, Prov. Cordillera, 5 km N of Emboscada, Rio
Piribebuy, forest with bromeliads, dead wood and leaves, 05.04.1985, leg. Expedition
zoologique du Museum de Geneve. - Paratype: 1 juv. (MHNG), same locality, together with
holotype.

Name: To emphasize the only slightly modified legs of the 6 and the relatively simple
gonopod structure.

Diagnosis: Differs from congeners by the relatively small size, the only slight-
ly modified legs of the c3, and the relatively simple solenophore structure.

Description: Length of holotype ca 12 mm, width of midbody metazona
1.5 mm. Coloration in alcohol uniformly light red-brown, anterior body portion

818

S. I. GOLOVATCH

Figs 26-32

Mestosorna simplex sp. n., 6 holotype: 26) anterior body portion, lateral view; 27) left halves of
segments 10 and 1 1 , dorsal view; 28) sternal structures between coxae 4 and 5 (coxae 4 on top);
29) leg 15; 30-32) right gonopod, medial, submedial and lateral views, respectively. Scale bar
0.6 (26-29) and 0.3 mm (30-32).

slightly more intensely reddish; penultimate antennomere darker brown, tip of antenna
pallid; legs paler, yellow-brown.

Body strongylosomoid, not moniliform, with poorly developed but evident
paraterga. Postcollar constriction apparent, width of head = collum > 2 = 4 « 5-16; on
segments 17-20 trunk gradually and gently tapering toward telson both in width and in
height. Antennae medium-sized, slightly clavate, in situ reaching beyond segment 2
dorsally (Fig. 26), a little shorter in juvenile. Paraterga 2 and 5-19 delimited by an
evident sulcus not only dorsally but also ventrocaudally (Fig. 26), all set low,
considerably larger on pore-bearing segments than on poreless ones (Fig. 27), only on
segments 18 and 19 slightly projecting beyond rear tergal contour. Tegument smooth
and shining. Limbus thin, caudal margin entire. Metaterga 5-17 with a faint but

PARADOXOSOMATIDAE IN PARAGUAY

819

apparent transverse sulcus far from reaching base of paratergum, the sulcus being very
slightly sinuate anteromedially; axial impression on metaterga interrupted in the mid-
dle, barely visible; tergal setae untraceable (Figs 26, 27). Stricture between pro- and
metazona thin and shallow (Figs 26, 27). Ozopores lateral, lying on paraterga at ca 1/4
metazonital length away from caudal edge (Figs 26, 27). Pleurostemal carinae like
narrow keels delimited by a sulcus dorsally, devoid of a caudal spinule (Fig. 26), trace-
able until segment 17, on further segments wanting. Epiproct rather long, coniform,
narrowly truncate at tip in dorsal view, in <5 a little longer than in juvenile. Hypoproct
roundly triangular, tip evidently rounded, straight; 1+1 paramedian setae at caudal edge
rather strongly separated from each other.

Sterna modestly setose, mainly unmodified but both with an evident, deeply
emarginate, setose, linguiform outgrowth directed anteroventrally between coxae 4 of
8 and with a paramedian pair of bunches of setae between coxae 5 of 8 (Fig. 28).

Legs only slightly modified, in 8 with dense tarsal brushes starting from leg-
pair 1 and with dense tibial brushes starting from leg-pair 2 (Fig. 29), both these
brushes thinning out toward telson but absent only from last two leg-pairs. Each coxa
2 of <5 with a low disto ventral cone bearing a gonopore.

Gonopods (Figs 30-32) relatively simple. Coxite subcylindrical, not particularly
elongate, sparsely setose disto ventrally. Telopodite subfalcate, a densely setose
prefemoral part normal in shape, much shorter than femorite. Solenophore unciform,
somewhat constricted at base, relatively stout and simple; lobe m at base of both
solenomere and lamina medialis highly inconspicuous, ledge-shaped, lamina lateralis
slightly better developed than lamina medialis, tips of both subtruncate.

Remarks: Due to the basally strongly constricted solenophore, M. simplex sp. n.
joins group III in the sense of Jeekel (1963), which has hitherto been known to
comprise only three species, one each from the Brazilian states of Pernambuco,
Alagoas and Sao Paulo. However, the new species differs in its smaller size, in the
absence of femoral and some sternal modifications in the 8 , and in the particularly
simple gonopod structure.

Mestosoma crassipes sp. n. Figs 33—42

Material: Holotype 8 (MHNG), Paraguay, Prov. Concepcion, Ao. Tagatya-mi, small
grove, near “gue”, sifted litter, 23.10.1985, leg. Expedition zoologique du Museum de Geneve.

Name: To emphasize the modified and inflated legs of the 8 .

Diagnosis: Differs from congeners by the relatively small size, the conspicu-
ously modified legs of the 8 , and by the relatively complex gonopod structure.

Description: Length of holotype ca 12 mm, width of midbody metazona
1 .3 mm. Coloration in alcohol uniformly light brown; antennae brown, penultimate an-
tennomere darker brown; tip of antenna pallid; legs slightly paler.

Body strongylosomoid, not moniliform, much like in M. simplex sp. n. Post-
collar constriction apparent, width of head ? collum =4>2=3<5< 6(7)- 16; on seg-
ments 17-20 trunk gradually and gently tapering toward telson both in width and in
height. Antennae medium-sized, slightly clavate, in situ reaching beyond segment 2
dorsally (Fig. 33). All paraterga set low, delimited by an evident sulcus not only
dorsally but also ventrally to ventrocaudally (Fig. 33), on segment 19 particularly flat

820

S. I. GOLOVATCH

Figs 33-42

Mestosoma crassipes sp. n., 6 holotype: 33) anterior body portion, lateral view; 34) left half of
metatergum 10, dorsal view; 35) caudal body portion, lateral view; 36) sternal structures be-
tween coxae 3 and 4 (coxae 3 on top); 37) leg 7; 38) leg 9; 39-42) left gonopod, submedial, sub-
lateral, ventrolateral and anterodorsal views, respectively. Scale bars 1 .0 (33-35), 0.5 (36-38) and
0.3 mm (39-42).

in lateral view, considerably larger on pore-bearing segments than on poreless ones
(Figs 33, 34), never projecting beyond rear tergal contour. Tegument smooth and shin-
ing. Limbus thin, caudal margin entire. Metaterga 5-18 with a faint but apparent trans-
verse sulcus far from reaching base of paratergum, the sulcus being slightly sinuate an-
teromedially; axial impression on metaterga wanting; tergal setae untraceable (Figs 33,
34). Stricture between pro- and metazona thin and shallow (Figs 33, 34). Ozopores lat-
eral, lying on paraterga at ca 1/4 metazonital length away from caudal edge (Figs 33,
34). Pleurotergal carinae like narrow keels delimited by a sulcus dorsally, devoid of a
caudal spinule (Fig. 33), traceable until segment 17, on following segments wanting.
Epiproct rather long (Fig. 35), coniform, narrowly truncate at tip in dorsal view.
Hypoproct triangular, tip pointed, straight; 1 + 1 paramedian setae at caudal edge widely
separated from each other (Fig. 35).

Sterna mainly modestly setose, postgonopodial ones unmodified, pregonopodi-
al ones with a paramedian pair of bunches of setae on small cones between coxae 3 and

PARADOXOSOMATIDAE IN PARAGUAY

821

4 of 8 (Fig. 36); sterna between coxae 5-7 of S concave and with some particularly
long setae (like in Fig. 36).

Legs modified, with dense tarsal brushes starting from leg-pair 1 and with dense
tibial brushes starting from leg-pair 2 (Figs 37, 38), both these brushes thinning out to-
ward telson and virtually absent only from last two leg-pairs. Femora 4-6 conspicu-
ously inflated and each with an evident distoventral tooth. Legs 7 (Fig. 37) with a dis-
toventral cone on each coxa and a little less strongly incrassate femora, the tooth situ-
ated more ventro-apically. Femora 9-11 (Fig. 38) nearly normal, ventral knobs in dis-
tal one- third gradually disappearing toward leg-pair 12. Each coxa 2 with a low dis-
toventral cone bearing a gonopore.

Gonopods (Figs 40-42) somewhat more complex than in the previous congener,
especially so due to a longitudinal sulcus on dorsal face of femorite, a digitiform
process (m), not a ledge-shaped structure lying at base of lamina lateralis, as well as a
short, unciform and pointed solenophore tip.

Remarks: This new species seems to be particularly close to M. alticola and to
a few other congeners possessing a small but evident process (m) at base of both the
solenophore and solenomere (see Attems, 1937). The colour pattern in M. crassipes sp.
n. resembles that of M. femorale , yet the gonopod structure is quite different (see
Schubart, 1943). Due to the presence of two small tubercles between coxae 3 of the 6
and an attenuated tip of the solenophore, the new species is similar to M. perfidum , but
the latter species is larger (17-18 mm long and 1. 9-2.0 mm wide versus 12 and
1.3 mm), and their gonopods are very different as well.

IDENTIFICATION KEYS

Key to tribes and genera of Paradoxosomatidae occurring in Paraguay and

ADJACENT AREAS

1 Paraterga strongly developed, wing-like. (Introduced, normally synan-

thropic species) 2

Paraterga poorly developed to missing (Figs 1, 10, 18, 26, 33) (Tribe
Catharosomatini, autochthonous species) 4

2 Metaterga densely papillate and setose; a bituberculate process between
coxae 4 of 8 ; femora 4-7 of S each with a ventro-parabasal knob; gono-
pod femorite stout, solenophore with a large basal prong ventrally

Tribe Sulciferini: Chondromorpha (C. xanthotricha)

Metaterga not papillate, at most poorly setose; sternum between coxae 4

of 6 without process; legs of 6 without such femoral tubercles; gono-
femorite not so stout, almost as long as to longer than solenophore 3

3 Caudal comers of paraterga surpassing rear tergal contour only on cau-
dalmost segments (16-19); gonofemorite relatively short, distinctly
broadened distally; solenophore bifid, at base with a long apical process

as well as a prong and a lobe ventrally . . Tribe Sulciferini: Oxidus ( O . gracilis)
Caudal comers of paraterga surpassing rear tergal contour on all body
segments; gonopod telopodite very slender and simple, rod-shaped; go-
nofemorite slightly longer than solenophore, latter without outgrowths

at base but with a minute lobule at tip

Tribe Orthomorphini: Orthomorpha ( O . coarctata)

822

4

5

6

7

8

9

10

11

12

S. I. GOLOVATCH

Gonopod devoid of a solenophore, solenomere free 5

Gonopod with a solenophore sheathing and supporting a flagelliform

solenomere 9

Gonopod telopodite extremely simple, subfalcate to subcircular, devoid
of any outgrowths at base of a subflagelliform solenomere . . . Gonodrepanum
Gonopod telopodite more complex, with 1-2 processes or dilatations in

femoral or postfemoral region 6

Gonofemorite with a large, medial, lobuliform dilatation; solenomere

with a small tooth near tip Mogyella (M. nano )

Gonofemorite without a large dilatation but sometimes with a tooth dis-
tally or parabasally; 1-2 dilatations or processes at base of solenomere,

latter sometimes stout and thick 7

Body small, 0.7 mm wide. Gonopod telopodite subfalcate, much like in
Gonodrepanum ; femorite with a small ventro-parabasal tooth; a small
but evident, elongated, subtriangular dilatation/lobe fused ventrally at

base with a flagelliform solenomere Pseudo gonodrepanum (P. scitum)

Body medium-sized, >1.0 mm wide. Gonopod telopodite mostly sub-
erect, usually with two independent teeth/outgrowths at base of soleno-
mere 8

Entire gonopod telopodite suberect, with an evident process at base of a
rather thick, sometimes bifid solenomere; a setose, linguiform, sternal

process between coxae 4 of 6 Habrodesmoides

Only gonopod femorite elongated and suberect; solenomere subflagelli-
and unciform, acuminate, at base with two relatively small teeth/out-
growths; sternal process between coxae 4 of 8 either missing or repre-
sented by two paramedian knobs Ologonosoma

Solenophore simple, subflagelliform, acuminate, supporting a similarly

flagelliform solenomere 10

Solenophore complex, sometimes with a process or lobe at base, usually
consisting of several more or less folded lobes/laminae, almost entirely

sheathing a flagelliform solenomere 11

Body width 0.7 mm. Pre femoral (setose) part of gonopod much shorter
than acropodite; femorite with a conspicuous lamella on medial side;
both solenophore and solenomere somewhat reduced, about half as long

as femorite Mogyosoma (M. hamatum)

Body width 1.1- 1.5 mm. Prefemoral portion of gonopod elongated,
nearly as long as acropodite; femorite without a lobe on medial side;

both solenophore and solenomere longer than femorite

Gonodrepanoides (G. travassosi)

Solenophore relatively simple but with a large lateral branch/process at

base Promestosoma (P. boggianii)

Solenophore more complex, usually consisting of several lobes but

devoid of a large lateral branch at base 12

Sternal cones usually present (Figs 4 & 15); each coxa 2 of 6 with a
strong disto ventral process carrying a gonopore (Figs 1 & 14); tibiae of

PARADOXOSOMATIDAE IN PARAGUAY

823

8 usually conspicuously inflated (Figs 5 & 16). Prefemoral (densely
setose) portion of gonopod hypertrophied, about as long as femorite

Catharosoma

Sternal cones usually absent; normally each coxa 2 of 8 with a small
cone carrying a gonopore; tibiae of 8 not inflated. Prefemoral portion of
gonopod normal, shorter than femorite 13

13 Tarsal brushes often absent even in 8 ; stemite between coxa 3 of 8 with
a conspicuous comb- or tongue-shaped structure (Fig. 20); stemite
between coxae 5 of 8 with a particularly large process directed antero-
ventrally and covering a boss or protuberance if any between coxae 4

Broelemannopus

Tarsal brushes in 8 mostly present; stemite between coxa 3 of 8 usually
without protuberances; a particularly large process directed anteroven-
trally and placed between coxae 4, not 5, of 8 Mestosoma

Key to Broelemannopus species of Paraguay and adjacent areas

1 Body of adult < 10 mm long and <1.0 mm wide; epiproct topped with
a prominent claw-shaped uncus directed slightly dorsad (Fig. 19); cones

absent at least between front coxae of postgonopodial sterna 2

Body > 14 mm long and > 1 .6 mm wide; epiproct without central uncus

at tip, latter usually bifid; sternal cones in 8 present between both pairs
of legs of each segment; Brazil 3

2 Lobe between coxae 4 and 5 of 8 linguiform, entire (Fig. 20); sternal
cones on midbody segments of 8 totally absent (Fig. 21); solenophore

lobe p small (Figs 23-25); Paraguay B. minutus

Lobe between coxae 4 and 5 of 8 divided distally; sternal cones in 8
present between posterior coxae of segments 8-18; solenophore lobe p
large; Sao Paulo State, Brazil B. ibitiensis

3 Body 1. 6-2.2 mm (<?) or 1. 6-2.4 mm wide (?). Paraterga traceable as
low arcuated sulci on segments 2-4, on following segments only as
slight impressions visible near caudal margin; pleurostemal carinae

traceable until segment 14 B . pirassunungensis

Body 2.3-3 .3 mm wide (8, 2). Each paratergite traceable as a low
arcuated crest at least on segment 2; pleurostemal carinae visible until
segment 15 4

4 Sternal lobe between coxae 3 of 8 comb-shaped, entire distally; coxae
2-6 of 8 each with a disto ventral coniform process; central protuberance
between coxae 7 of 8 absent; distal end of solenophore not divided

B. escaramncensis

Sternal lobe between coxae 3 of 8 linguiform, deeply incised distally;

only coxae 2 and 3 of 8 each with a coniform process; central protu-
berance between coxae 7 of 8 present; distal end of solenophore deeply
divided into two large lobes, p being one of these B. glabratus

824

S. I. GOLOVATCH

Key to Catharosoma species

1

2

3

4

5

6

7

8

9

10

Sternum between coxae 6 of 6 with a single, undivided process or

swelling 2

Sternum between coxae 6 of S either with a deeply divided process or
with two distinct paramedian processes, or with two paramedian knobs

with bunches of long setae 5

Hypoproct tip unciform; gonopod femorite with a prominent, subtrian-

gular, distal lobe C. digitale

Hypoproct tip straight; gonofemorite without a prominent, subtrian-

gular, distal lobe 3

Body width about 2.2 mm; no sternal modifications except pilosity be-
tween coxae 5 of 6 ; tip of solenophore simple, like a broad and sub-
acuminate lobe C. mixtum

Body width 3.0-3 .2 mm; a swelling and/or a paramedian pair of knobs

between coxae 5 of $ ; tip of solenophore more complex 4

Paraterga 2 like swellings, paraterga 3 and 4 sulciform; light subtri an-
gular spots on posterior parts of proterga and on anterior parts of
metaterga against a dark background; apical piece of solenophore short,

acuminate C. peraccae

Paraterga 2-4 like low crests; background coloration pale yellowish with
a wide castaneous axial stripe; apex of solenophore very broad and of

rather irregular shape C. mesorphinum

Even paraterga 2 expressed as sulci; no sternal modifications except

pilosity between coxae 5 of 6 C. myrmekurum

Crest-like paraterga present at least on segment 2 (Figs 1 & 10), often
ridge/crest-like even on segments 2-4; sternal modifications present

between coxae 5 of d 6

Paraterga 2-4 distinctly crest-like, onward sulciform on segments 5,7,9
and 10; a distinct bilobed process present between coxae 3 of 6 C.palmatum
Paraterga even more poorly developed, totally untraceable on segments
behind 4 th 7

Body about 12 mm long and 1.3 mm wide; coxae 2 of 6 produced
distally into a small mammiform process; gonofemorite with a medio-

parabasal digitiform process C. curitibense

Body width > 1 .7 mm; coxae 2 of 6 usually produced into a more or less

prominent process (Fig. 14); gonofemorite devoid of such a process 8

Hypoproct unciform, its tip directed ventrad 9

Hypoproct tip straight 12

Dorsum rather dark, red-brown, with two yellowish paramedian stripes;

pleurostemal carinae visible only until segment 15 C. taeniatum

Dorsum pale but with two dark, brownish paramedian stripes; pleu-
rostemal carinae visible at least until segment 16 10

Body of 6 1. 9-2.1 mm wide; a large and deeply incised process present
between coxae 6 of 6 (Fig. 3) C. bilineatum

PARADOXOSOMATIDAE IN PARAGUAY

825

Body width >2.6 mm; only a pair of small paramedian knobs between
coxae 6 of S 11

11 Apical papillae on epiproct acuminate, elongate and directed distodor-

sad; gonofemorite strongly broadened distad C. mesoxanthum

Apical papillae on epiproct small, simple, inconspicuous; gonofemorite
slender C. intermedium

12 A paramedian pair of low setigerous crests present between coxae 3 of

S\ gonopod telopodite elongate and circular (Fig. 17) C. mahnerti

At most a paramedian pair of bunches of setae present between coxae 3

of S ; gonopod telopodite falcate and stout 13

13 A single protuberance between coxae 5 of S and a paramedian pair of

conical processes between coxae 6 of 6 C. palustre

Two setigerous knobs on a swelling between coxae 5 of 6 and either a
swelling or a paramedian pair of spinules between coxae 6 of 6 14

14 Body width 2.0-2.3 (<?) to 2.5 mm (2); lobe p of solenophore ancori-

form and bifid C. paraguay ense

Body width 2.6 mm (<3); tip of lobe p of solenophore blunt, devoid of
teeth C. hoffinani

Key to Gonodrepanum species

1 Coloration of adults normally castaneous brown with a wide, uninter-
rupted, axial, contrastingly creamy stripe; Rio de Janeiro, Brazil 2

Coloration uniformly castaneous to black, usually devoid of a contrast-
ingly pallid stripe, or axial stripe/line thin and interrupted; Argentina
and/or Brazil 3

2 Smaller, only known from 2: 16-18 mm long and 1. 8-2.0 mm wide;

clear sulci in place of paraterga 2-4 G.flavolineatum

2 18-21 mm long and 2. 2-2. 4 mm wide, <? 15-16 mm long and 1 .5-1 .6

mm wide; paraterga on segment 2 like crests, on segments 3 and 4 like
clear sulci, on following segments like striae; paramedian pairs of
setigerous protuberances present between coxae 3-5 of 8 G. torresae

3 Body particularly slender: S 20 mm long and 1 .2 mm wide. Dorsum and
sides uniformly blackish; paraterga 2-4 crest-like, on following

segments sulciform; sternal cones absent; Brazil and Argentina

G. drepanephoron

Body neither so long (< 14 mm even in 2 ) nor so slender. Coloration of

adults usually castaneous, sometimes yellowish; paraterga represented

by sulci at most; sternal cones present; Brazil 4

4 Epiproct topped with two long, claw- shaped, diverging unci directed

dorsocaudad; a pale axial stripe/line absent 5

Epiproct without such unci; a pale, axial, interrupted stripe/line usually
present 6

Body 12-14 mm long and 1.3- 1.7 mm wide; a small ventral tubercle
present only on femora 3 and 4 of <? ; sternal protuberance between coxae

826

S. I. GOLOVATCH

3 of S bifid, sterna between coxae 4 and 5 devoid of protuberances;

solenomere subcircular G. grajahuense

Body 9-12 mm long and 1.0-1. 3 mm wide; a distoventral tubercle
present/traceable on femora 3-9 of c J; median sternal protuberances
present between coxae 3-5 of $ ; solenomere regularly and gently sub-
falcate G. furcation

6 Body about 14 mm long and 1.3- 1.5 mm wide. Metatergal sulci absent;
femora of 6 not modified; a large median process with two knobs
behind and between coxae 4 of 6 ; solenomere relatively short and

suberect; Santa Catarina, Brazil G. levisetum

(G. levisetum var. coniferum seems to only represent a colour form dis-
tinguished by a yellowish body devoid of a pattern)

Body 11-12.5 mm long and 1.1 mm wide. Metatergal sulci light but
present; a pair of diverging processes between coxae 5 of 6 ; solenomere
relatively long, subcircular; Rio de Janeiro, Brazil G.falciferum

Key to Habrodesmoides species of Paraguay and adjacent areas

1 Paraterga replaced by a low arcuated ridge on segment 2, by sulci on
segments 3 and 4; pleurostemal carinae traceable until segment 18; a
small protuberance present between coxae 5 of 8 ; solenomere retrorse,

with a slender, subfalcate, simple process at base H. costalimai

Segments 2-18 with only slight sulci/striae in place of paraterga; pleuro-
stemal carinae present on segments 2-7; stemite between coxae 5 of S
devoid of a protuberance; solenomere directed distoventrad, process at
its base massive, stout and branched H. perturbans

Key to Mestosoma species of Paraguay and adjacent areas

1 Sterna between coxae (3)4-6(7) of $ each with a more or less distinct

median process or protuberance; sternal cones in 6 absent 2

At least some of these sterna devoid of a protuberance; sternal cones in

S often present 5

2 Body width > 2.3 mm; metatergal sulcus present on segments subse-
quent to 5 th ; no single sternal process between coxae 6 and 7 of 6 3

Body width < 1 .5 mm (<3); metatergal sulci absent; a single, prominent,
sternal process between coxae 6 and 7 of 6 ; Bolivia 4

3 Coloration uniformly brownish; sternal processes or protuberances

present between coxae 3-6 of S ; Rio de Janeiro, Brazil M. carioca

With a wide pale axial stripe against a dark olive-brown background;
sternal process present between coxae 4 of 6, only protuberances or
swellings between coxae 5 and 6 in 6 ; Bolivia M . schindleri

4 Body width 1.0 mm (c?); colour pattern in adults indistinct, body uni-

formly yellowish; femora 3-5 of c 3 each with a ventral tubercle . . . M. alticola
Body width about 1.5 mm (cJ); a wide, pale, axial stripe against a cho-
colate-brown background; femora in i apparently without ventral
tubercles M. boliviae

5

6

7

8

9

10

11

12

13

14

15

PARADOXOSOMATIDAE IN PARAGUAY

827

Sterna of segment 6 in 3 with a pair of processes at anterior border,
sterna of segment 5 of 3 devoid of processes; neither tibial nor tarsal

brushes present M. differens

Sterna of segment 6 in 3 without processes, those of segment 5 of 3
with or without processes; tibial and tarsal brushes in 3 normally present ... 6
Colour pattern relatively distinct, usually entire dorsum or axial stripe
pale to yellowish, sometimes this stripe divided into two paramedian
stripes or broken into series of spots, more or less strongly contrasting

with a dark, normally brown background coloration of the sides 7

Colour pattern indistinct, normally dorsum and sides uniformly brown .... 14

Sternal process between coxae 4 of <5 present, conspicuous 8

Sternal process between coxae 4 of c ? absent to very small 10

Body width < 2.0 mm. Dorsum with axial stripe(s), background
coloration brown; gonofemorite not very broad, solenophore evidently

longer than 1/2 femorite 9

Body width >2.0 mm. Dorsum with an axial series of yellowish spots,
background coloration dark (red-)brown to black; gonofemorite
conspicuously broadened, solenophore scarcely half as long as femorite;

Paraguay 10

Dorsum with two yellowish paramedian stripes; tibial and tarsal brushes
in 3 present; a bilobate process present between coxae 4 of c 3 ; Paraguay

M . vittatum

Dorsum with two series of paramedian spots; tibial and tarsal brushes in
<3 absent; a lobe-shaped process present between coxae 4 of 3 ; Bolivia

M. montanum

Solenophore with 2-3 prongs/spines subapically 11

End of solenophore subtriangular, often rounded 12

Legs uniformly blackish; a pair of paramedian knobs between coxae 3
of 3 ; femora 4-7 in 3 each with a ventral tubercle, each coxa 7 of 3

with a distoventral process M. tricuspis

Legs brown; a process present between coxae 4 of 3 ; apparently neither
femora nor coxae of 3 with ventral tubercles/processes . . M. pseudomorphum
Body width about 3.0 mm. Dorsum with a pale, sometimes yellowish
axial stripe; solenophore strongly curved proximad, its end almost in

contact with base of femorite M. salvadorii

Body width about 2.0 mm. Dorsum with an axial series of conspicuous

yellowish spots; solenophore not so strongly curved 13

Median spots situated on proterga and in anterior parts of metaterga;
solenophore in medial view almost twice as slender as distofemoral part

M. camercini

Median spots lying only in middle of metaterga; solenophore in medial

view nearly as broad as distofemoral part M . kcilliston

A single process present between coxae 4 of 3 15

Either two cones/tubercles or nothing between coxae 4 in 3 17

Body width about 2.0 mm. Sternal process between coxae 4 of 3 short
and subquadrate; solenophore bifid apically M. bcilzanii

828

S. I. GOLOVATCH

Body width 1 .3-1 .8 mm. Shape of sternal process between coxae 4 in d
and of solenophore different 16

16 Metatergal sulci distinct on segment 5 and following segments; sternal
process between coxae 4 of d short and conical; gonofemorite much

broader than solenophore M. lugubre

Metatergal sulci absent; sternal process between coxae 4 of d long and
rectangular; gonofemorite considerably slenderer than solenophore M. borellii

17 Solenophore base conspicuously constricted (Figs 30-32) M. simplex

Solenophore base broader 18

18 Body length about 12 mm, width 1 .3 mm. Metatergal sulcus visible also
on segment 18; pleurostemal carinae present until segment 17; a disto-
ventral tubercle present on femora 4-12 of d , and a process on coxae 9
of d (Fig. 38); a characteristic process (m) at base of solenophore, tip

of solenophore pointed (Figs 39-42) M. crassipes

Body length at least 17 mm, width 1.8 mm. Metatergal sulci either
absent or at most traceable only until segment 17; distofemoral tubercles
present until leg 10 of d at most; coxae 9 of d normal; no process at
base of solenophore, tip of solenophore more or less rounded 19

19 Body length 30-34 mm, width 3. 8-4 .2 mm; tip of solenophore very

broadly subtruncate M. truncatum

Body length < 29 mm, width <3.5 mm; tip of solenomere not subtruncate . 20

20 Faint sternal cones traceable between coxae 3 and behind segment 7 of

d ; dis to ventral tubercles present only on femora 9 and 10 of d . . M.perfidum
Noteworthy sternal modifications absent; distoventral tubercles usually
present only until femora 7 of d 21

21 Tip of solenophore bilobate, rather deeply emarginate/notched in the

middle, forming more or less equal, rounded lobes 22

Tip of solenophore different 23

22 Coloration blackish, legs red-brown; both lobes of solenophore tip sub-
equal; Bolivia M. dereliction

Coloration dark brown, legs light brown; proximal lobe of solenophore

tip somewhat smaller than distal one; Paraguay M. pulvillatum

23 Tip of solenophore subacuminate, narrowly rounded; Bolivia . . . M. luctuosum

Tip of solenophore broadly rounded; Brazil and/or Paraguay 4

24 Body coloration dark, legs and venter contrastingly pale; pleurostemal
carinae traceable until segment 15; a distoventral tubercle present only

on femora 7 of d ; ventral brushes on tibiae of d absent M. bicolor

Body coloration brown, legs and venter only slightly paler than remain-
ing body; pleurostemal carinae traceable until segment 16; distoventral
tubercles present on femora 4-7 of d ; ventral brushes present both on
tibiae and tarsi in d M.femorale

Key to Ologonosoma species

1 Sternal process between coxae 4 of d missing; solenomere retrorse;

Brazil (Parana) O. iguassuense

PARADOXOSOMATIDAE IN PARAGUAY

829

Sternal process between coxae 4 of & represented by two paramedian
knobs; solenomere directed distoventrad; Paraguay O. sanctum

ACKNOWLEDGEMENTS

I am most grateful to Dr Volker Mahnert (MHNG), who entrusted me this valu-
able material for study. Peter Schwendinger (MHNG) kindly edited an advanced draft.

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